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Thursday, September 8, 2016

ASHG 2016 abstracts

The American Society of Human Genetics (ASHG) 2016 Meeting Online Planner and Abstract Search can be accessed here. Unless I'm missing something, there's surprisingly little ancient genomics stuff this year considering the huge advances in this area recently.

However, Iain Mathieson has an interesting contribution about interactions between Balkan foragers and early Balkan farmers. Reading between the lines, I reckon we can probably expect some R1b in these Balkan ancients. This shouldn't be surprising, however, considering the publication earlier this year of the Ice Age Villabruna genome from present-day northeast Italy, which belonged to R1b and, as pointed out by myself, also showed strong genome-wide ties to ancient Siberians (see here).

Oleg Balanovsky has been acting a little strange lately; seemingly going out of his way to fight the Kurgan Indo-European paradigm in various papers and talks. In this poster he underlines the fact that the Yamnaya people from the North Caspian region did not totally vacate their homeland nor moved en masse to Western Europe, but actually left most of their descendants in the North Caspian. OK Oleg, but almost all modern day Europeans, especially those from across Northern and Eastern Europe, do derive a lot of their ancestry from Bronze Age steppe groups closely related to the Yamnaya people of the North Caspian. This will become even more obvious soon with the publication of new ancient data from the Baltic region.

And here we have what appears to be yet another hilariously wrong paper in the making on the population history of South Asia. Why do they mention Anatolia? How about the Arctic Circle? Enough already. Get some goddamn ancient genomes from South Asia and finally do it right.

236 comments:

Using whole-genome sequencing to shed insight on the complex prehistory of Sardinia

"In unsupervised analyses, a Sardinian-specific ancestry component correlates with shared drift parameters with both the Neolithic farmers and Paleolithic hunter-gatherers, and is consistent with supervised estimates of ancestry proportions in which the Ogliastra individuals have higher Neolithic farmer and Paleolithic hunter-gatherer ancestries, while individuals from the rest of the island show an infusion of the pastoralist ancestry."

I think the Balanovsky paper looks very interesting. Here is its abstract :-

"For haplogroup R1b, we identified a previously unstudied “eastern” branch, R1b-GG400, found in East Europeans and West Asians and forming a brother clade to the “western” branch R1b-L51 found in West Europeans. The ancient samples from the Yamnaya archaeological culture are located on this eastern branch, showing that the paternal descendants of the Yamnaya population – in contrast to the published autosomal findings - still live in the Pontic steppe and were not an important source of paternal lineages in present-day West Europeans."

This should make Gioiello quite happy I presume. If Yamnaya did not contribute R1b in West Europeans, how did it manage to have such a massive autosomal impact in Europe, as per Haak et al ? This is quite intriguing.

Also, The Balanovsky paper seems to support the Iranian/Central Asian link to Yamnaya, which Lazaridis et al proposed. Here we may also take note of the Indian/Iranian wolf & dog ancestry in the ancient dog from a corded ware site in Germany.

It may also be noted that Lactase Persistence (LP) only appears in Europe during the Bronze Age and quite possibly from the steppe. But earlier to the steppe, as per the Broushaki et al paper, LP was perhaps already present in Iranian Neolithic samples.

MfA: Maps of effective migration as a summary of human genetic diversity.https://ep70.eventpilot.us/web/page.php?page=IntHtml&project=ASHG16&id=160122526

Comment from them that stood out to me:

"We apply EEMS on global, continental, and sub-continental scales, analyzing genetic data from 8,740 individuals from 469 geographically localized populations, obtained from 24 different source studies. In addition to the major, well-known barriers such as the Sahara, Himalayas and Mediterranean, we detect barriers that correlate with historic language group boundaries (boundaries of Slavic and Bantu speakers with their neighbors), mountain ranges (Zagros, Caucasus, Ural) and marine features (English Channel, Adriatic Sea, Wallace line). We also identify regions showing high connectivity despite having geographic separation (Britain and Scandanavia, Iceland and Denmark, among the Lesser Sunda Islands). "

Some use of the EEMS maps they describe, for Europe only, in this recent one by two of the same authors- http://biorxiv.org/content/early/2016/09/04/073221 - "Recent advances in the study of fine-scale population structure in humans" on page 15.

Screenshot - http://imgur.com/a/Av1TO.

Also includes an interesting use of SpaceMix on only modern European samples, which provides a really good fit to geography. A very impressive fit if all that was used was purely the genetic data alone, better than PCA typically gives.

Another example of the EEMS methodology here, by Mathieson: http://mathii.github.io/review/2016/01/25/spatial-genetics-1 looks like with slightly different estimates of rates. There is also a SpaceMix here. The positions of recently admixed populations and the Europeans from one another vary from dimensions 1-2 of PCA.

Despite the recent surge of interest in ancient genomes, we show that there is still much to be elucidated about human demography from contemporary genomes. Here, we demonstrate the use of genealogical data to generate demographic insights from analysis of a large-scale, heterogeneous genetic data set. Specifically, we show that an unsupervised ADMIXTURE analysis of genotypes from 131,293 primarily US-born individuals, followed by a simple statistical analysis of the 3 million pedigree records linked to these genotype samples, yields novel insights into European genetic diversity ... Despite this, our ADMIXTURE analysis with K = 12 ancestral populations identifies 5 stable, genetically differentiated groups within Europe (with putative historical counterparts in parentheses): Ashkenazi Jewish, Irish (Celts), Eastern Europeans (Slavs), Scandinavians (Nordics) and Iberians, featuring Basques and Sardinians."

Sounds like advertising ;) with a naive methodology. Maybe they have found some new interesting trends with their very large sample though.

Davidski, you are a clever person and you won't be remembered amongst who banned me (except some sospensions), but, please, stay afar from the English Lady who put Villabruna in Trentino-Sud Tyrol. Villabruna isn't at the Italian/Slovenian border, firstly because it is in Belluno Province, Veneto, and Slovenians came there only after the fall of the Roman Empire.Balanosky isn't a little: he, and his alii, has demonstrated what I am saying from ten years. I save you and your blog the ferocious letter I posted on my FB page and sent all my friends privately.

I can be your friend but not FB sorry :( . I don't see the reason of not posting the letter here . I recognize you as a veteran geneticist . But of course Me, Jaydeep and Parasar are all different folks ( Marnie probably is copying this conversation right now ;) .

We have only results from the eastern parts of the Yamnaya culture. The Yamnaya is a large region. We don't have results from western parts of the Yamnaya culture yet but i am convinced that R1b-L51 will be found there.

What Tesmos writes ("We have only results from the eastern parts of the Yamnaya culture. The Yamnaya is a large region. We don't have results from western parts of the Yamnaya culture yet but i am convinced that R1b-L51 will be found there") make me be prudent yet. I spoke of tumuli digged in Middle East with corpses of many geneticists, above all with a PhD at Stanford or Harvard, and also old Italians that I named Cavallo Sforzesco, Farfugliani, Sgaramello. But the worst was for the blunders in service graves... Let's wait that also Western Yamnanya is digged, but perhaps all those Russians will know the difference between East and West? What do you think?

"There aren't any real links between the European steppe and Iran before the Bronze Age. The uniparental markers in these two regions pre-Bronze Age are from different galaxies."

Western Iran chalc are not ancestors of yamnaya. It would not make sense if they were since the regions are not contiguous.They have ancestral contributions from both west(anatolia, caucasus) and east( unsampled) that them a "good fit", thats all.

neither are yamnaya ancestors of south asia. They are not neighbors by any stretch. There a partial overlap in ADNA.

About the worst things is always better the digest (I gave you above) than the whole. But as a poet, and not a PhD in genetics, I give you an Italian verse of a Nobel Prize:"ma in attendere è gioia più compìta!"

I am already dealing with a person who says who knows things but is acting selfish , the last thing that I need is poetry , though our poetic tradition is ~4000 years old , but for now, I would love to read a clinical criticism instead of poetry! .

Where are the Bronze and Iron age Greek-Balkan genomes as well as the Maikop culture ancient genomes you talked about last year that were supposed to be released this year? Those are the ones i want to see most. Though the Baltic one looks interesting.

Sure. R1b1 came from the Italian refugee as HG but by 8000BC were part of Anatolia (south eastern) and learning how to become farmers and going heavy on goats and cattle domestication. And actually M269 was born when they moved to south Caucasus by 6000BC and lived quite happily until 4.900 BC and they started to flee and already had L23 as they flee North to the Kuban river (later Maykop land) and later kicked-out or moved as the “revolution” of pre great-civilizations was going still (leyletepe, to full Uruk) and the Cucuteni-Trypillian moved east to make the Maykop itself. So always the underdog moved up the samara river and there they were as Z2013 and then just lost themselves in the fog of time moving east (not West for sure).The L23 that move back to Eastern Anatolia by 4900 BC also ended up in Lower Egypt at the Delta of the Nile by 4800 bc (or 4500bc If were just the El Omari group) and those that moved West had L51 and enter Iberia from north Africa. Finally those that from lower Egypt moved north naturally follow kinship and ended up back in Georgia as the Colchis and Iberians (the Georgia ones) that the ancient Greeks always referred as Egyptians… Yeah, follow kinship, follow kinship.

Re: EEMS and barriers, that they are limited by recent migrations would be an issue - where you have two populations who are dissimilar to one another, and it's because one (or both) has recently migrated to where it is, then seems like the EEMS method will find a barrier.

Of course, it's true that there still is a barrier to migration, but not necessarily in the sense of "Isolation by distance dynamics rules, but in some regions it's relatively slower because of a barrier".

Method also seems super sensitive to sample set difference though in any case:

http://imgur.com/a/cjQb5

Clear differences between Mathieson using on Lazaridis 2014 dataset (which is lacking in Central European / Northwest European and Northern British Isles references) against Novembre's POPres (lacking in Eastern European sample sizes, where it has them sampled at all).

Mathieson's version creates more barriers in Southern France and Central Europe, e.g. south Germany and across the Pyrenees, but the POPRes data suggest there isn't a barrier there. Similarly inconsistencies between whether the English Channel and North Sea are barriers or highways...

(Busby's dataset which is a bit more evenly distributed across Europe than Human Origins 2013 - https://data.mendeley.com/datasets/ckz9mtgrjj/1 or the revised Human Origins from 2016 supplemented with a few more Northern and Western populations might be more useful for this, though both still lacking in Central Europe).

As a method also hard to apply to geographically overlapping populations with structured differences (e.g. divergent subsistance / caste populations in the same region).

@TesmosWe have only results from the eastern parts of the Yamnaya culture. The Yamnaya is a large region. We don't have results from western parts of the Yamnaya culture yet but i am convinced that R1b-L51 will be found there.

You’ll be badly disappointed no R1b-L51 folks will be found in western part of Yamnaya culture. This subclade appeared somewhere in Anatolia. Pontic steppe region during Sredny Stog – Yamnaya (western part) period was populated predominantly by carriers of subclade R1a-YP1272.

PgmNr 1162/W: Personal ancestry inference at the finest scale reveals more sub-structure in the UK.

Chromosome Painting has revealed genetic differences within the UK at a very fine scale [1], with structured genetic variation within a single county in some cases (such as Cornwall & South Wales). However, in that work, it was not possible to genetically distinguish much of England, which appeared as a single homogeneous group. Here, we describe an extension to the FineSTRUCTURE [2] clustering that can further distinguish ancestry even within England; for example, identifying regions such as Norfolk, the Midlands and the South as genetically distinct. The approach works by using the known county locations to craft genetic features to use in unsupervised clustering. Specifically, we group individuals by their geographic sampling location into reference donor populations. This forms an ancestry profile - which can be viewed as a careful choice of feature vector - that still allows unsupervised genetic clustering for all individuals... This also allows us to characterise the genetic relationship between the inferred clusters, several of which represent drift that is most strongly represented by a particular geographical region (including Cornwall, Wales, Scotland and the North of England) and others of which represent characteristic admixture proportions between these ancestral drifted populations.

Not as nice as apriori clustering without geographical information, but still interesting that this kind of high structuring is possible at all.

The number of allele counts was the highest in YRI and the lowest in CEU. Of particular interest, the mean of the tanning ability score is lower (i.e., lighter skin color) in Tongan (Polynesian) and Rawaki (Micronesia) populations than in HapMap-JPT and CHB (East Asian) populations. The present results suggest that the genetic variation that produces darker skin color of Oceanians is different from that of Asians.

Two Oceanian populations (Tongans / Rawaki) seem likely to share "tan reducing" alleles with Europeans than Asians are (even though Asians actually would probably have lower tanning than them). Fits with these references being likely to have some European ancestry.

Bell beaker was 50% Yamnaya-like not 50% Anatolia Chalcolithic-like. R1b-L51 is from Yamnya or at least the same region. Why can't some of you understand the dye is cast. Its been caste for a year now. R1b-m269 and r1a-m417 are from steppe, not Spain or turkey or wherever.

@Samuel AndrewsBell beaker was 50% Yamnaya-like not 50% Anatolia Chalcolithic-like. R1b-L51 is from Yamnya or at least the same region. Why can't some of you understand the dye is cast. Its been caste for a year now. R1b-m269 and r1a-m417 are from steppe, not Spain or turkey or wherever.

Anatolia was still badly populated territory at those times so small groups of R1b-L51 folks could migrate via Anatolia without even getting in contact with local people. This obsession with steppe homeland is so ridiculous. It’s based absolutely on nothing. There’s no any evidence but you still keep repeating this outrageous bullshit about steppe homeland. Both R1a and R1b folks came from the Iranian Plateau and not from steppe, India, Italia or wherever.

"please, stay afar from the English Lady who put Villabruna in Trentino-Sud Tyrol. Villabruna isn't at the Italian/Slovenian border, firstly because it is in Belluno Province, Veneto"

My apologies. That error was due to me blindly following Di Benedetto 2000, was said his samples were from Trentino-South Tyrol, though I should have realised from the map that Villabruna was just outside the border of that region. Of course Fu 2016 said where it was, but I did not notice. Shocking! It is now fixed.

@Azarov,"Anatolia was still badly populated territory at those times so small groups of R1b-L51 folks could migrate via Anatolia without even getting in contact with local people."

This isn't an explanation for how R1b-L51 traveled from Anatolia to West Europe without Bell Beaker Germans receiving Anatolia Chalcolithic-like ancestry, because German Bell Beaker doesn't have anytype of Chalcolithic West Asian ancestry. They had 50% Steppe ancestry because R1b-L51 came from the Steppe.

"This obsession with steppe homeland is so ridiculous. It’s based absolutely on nothing. There’s no any evidence but you still keep repeating this outrageous bullshit about steppe homeland. Both R1a and R1b folks came from the Iranian Plateau and not from steppe, India, Italia or wherever."

I know some think Steppe people were special and superior. I don't. I state the fact they migrated to various locations and contributed ancestry to various modern people. You and some others have an anti-Steppe bias because I guess you like to be rebels. Others I've read are either Kurd supremecists or the one Basque supremecist Maju. It ticks me off when after a year the Steppe origin of R1b-M269/R1a-M417 were proven, you guys keep saying bull shit about R1b and R1a being from Spain or Iran or Anatolia or any location on the planet you can find that isn't the Pontic Caspien Steppe.

@Olyumpus,"What bell beakers were 50% yamnaya? Not the "half breds" in Germany that we know were found 100 miles from the Bohemia BB group that we know were the sons/grandsons of CWC women, right?"

Just about all of them were half Yamnaya. Either keep up with the facts or don't post here. They have no genealogical records, so we wouldn't know if they were sons/grandsons of CWC women. My bet is they have little to do with CWC, that they are an independent Steppe/EEF/WHG mix.

The CWC mtDNA in Bell Beaker thing is a theory created by people who want R1b-L51 to be from anywhere from the Steppe, so their excuse for 50% Steppe ancestry in Bell Beaker is that they have CWC mtDNA and EEF/WHG Iberian Y DNA.

@Samuel Andrews,We are back to forgetting even those "half breeds" some call German BB (which some dispute if they were BB or we should call it something undermined) actually showed a fairly large Iberian Chalcolithic component (seen by whiffs of SSA)?

So those guys being loaded up with CWC dna, they should show some "Yamnaya" dna. Is anyone disputing CWC had relation with CWC? - No. So dont mix the subjects.

@Samuel Andrews,"no dna for..."We have better! - We have Nom Metric Dental traits that is a very very good proxy for Population DNA. And the "better" part is that I can sample 3 guys for full genome and they turn out to be the sons a gang raped girl snatched form 1000 miles way!

With Nm Dental trait (and we have the best ones done on Bell beakers) we are talking about 3000 samples from hundreds of diferent sites all over a vast geographical region.

What Nm dental says? that bell beakers (in portugal, spain, south france, north Italy, switerland and evem the Cspel group in Hungary are a tight Born Iberian group that did not mixed with locals.With Nm dental we also know that the boehmia group (all those german BB you guys love to bring about in here) where the only OTHER BB group that had contribution from local population. The only other. And that contribution was by CWC populations. Even better. Nm dental shows that BB and CWC exchanged women but men were very closed isolated.

Then it also shows a funny thing that only CWC men made a linkage from those groups and Unetice later groups. Not even CWC womem...

So, when talking about all this, lets always have these notions in mind, shall we!

"R1b1 came from the Italian refugee"More and more likely, not only for all what I have written and documented in these last ten years but above all for Villabruna 14000 ya.

"but by 8000BC were part of Anatolia (south eastern)"Which proof about that? Of course Villabruna migrated to Anatolia and Middle East (beyond Samara), but it isn't said that they were hg. Y R1b1. Italy had also certainly hg. I, but I think also J, some subclades of G, we don't know about H2 present in Sardinia and Iberia (we don't know if it came from Anatolia to Italy or before from Italyto Anatolia).So far no R1b has been found in Anatolia nor in Middle East (Natufians), neither J if not in recent time (less than 5000 years), and certainly Italy has older subclades, even though the expansion happened very likely from Caucasus."And actually M269 was born when they moved to south Caucasus by 6000BC"Which proofs about that? R-M269-PF7562 has been found around the Black Sea, but the huge subclade R-PF7563 is overwhelmingly Italian and Western European.All the rest you say is proofs-free, whereas I have published tons of documents explaining why also the R-L23 subclades of Western Europe don't derive from Samara and as this paper of Balanovsky et al. seems to demonstrate.R1b1a2-L23-Z2110* with more than 90 private SNP is also my haplotype surely in Western Europe from at least 6000 years as to YFull, at least 7200 as to me.

That abstract about the early Balkan farmers is interesting. But as always a bit cryptic:

"admixture between the regions first farmers and its indigenous hunter-gatherers was complex, with evidence of local admixture from hunter-gatherers related to those from Scandinavia and Eastern Europe."

And:

"The hunter-gatherer admixture in the early farmers of the Balkans is not closely related to the hunter-gatherer admixture that is predominant in present-day Europeans."

How to make those two sentences compatible? The first ones quite clearly implies they had ANE. The second one might imply that they belonged to a nowadays minor Y-Chromosome lineage (not I2a, R1a or R1b)?

In any case:

"The population admixture was patchy across both space and time, varying in magnitude between 0% and 30% for different early Balkan farming populations."

This is something genuinely new and important. If Neolithic Farmers from around Bulgaria (that seems to be from where the samples might be from) had up to 30% SHG/EHG admixture, that would have significant consequences in our understanding of LN/BA populations.

If Neolithic Farmers from around Bulgaria (that seems to be from where the samples might be from) had up to 30% SHG/EHG admixture, that would have significant consequences in our understanding of LN/BA populations.

It doesn't change any of the main models.

EHG admixture in modern Europeans is entirely from the steppe, not the Balkans or Northeastern Europe.

It does make sense in light of what we have with IRC KO1 having an excess of sharing with SHG relative to what would be expected from other members of the WHG clade.

But yeah, hard to see what they mean by that second quote, and why it shouldn't have some effect on the overall model.

If you're measuring the impact of Yamnaya, as EHG+CHG, via the ANE shift (which with some extra detail is how these formal models work), and then you have a population which is SHG+AN or EHG+AN, then that population should lead to some difference in estimates from modeling with EHG+CHG vs AN+WHG (MN_Chl West Europe).

That might make the estimates based on locating the CHG fraction of European ancestry and then extrapolating from that to Yamnaya impact more viable (e.g, Cassidy 2016 where the Bronze Age Rathlin was estimated via 40% CHG ADMIXTURE in Yamnaya "related to 6–13% in Rathlin ... implying a total of 14–33% Yamnaya ancestry and therefore 67–86% MN in the Irish Bronze Age").

But in balance also could make Western Yamnaya samples and a time series of these more important (if they were had a more AN+EHG and less of a EHG+CHG balance compared to the Yamnaya sample we know).

(Unless the Balkan farming populations just did not contribute at all!)

Corded Ware and Yamnaya are from the steppe, and present-day Europeans basically get their ANE/EHG shift from Corded Ware and Yamnaya. So they get it from the steppe. There's no doubt about that now, and if anyone still has any doubts then just wait a little.

Now, these Balkan farmers with their unusual EHG/SHG-like forager admixture may have contributed something to present-day Balkan populations, but unless they had a significant impact on the steppe ancestors of Corded Ware and Yamnaya, then they didn't contribute much to other present-day Europeans.

''You’ll be badly disappointed no R1b-L51 folks will be found in western part of Yamnaya culture. This subclade appeared somewhere in Anatolia. Pontic steppe region during Sredny Stog – Yamnaya (western part) period was populated predominantly by carriers of subclade R1a-YP1272. ''

Do you have evidence or serious clues that R1b-L51 appeared somewhere in Anatolia? It doesn't make sense that the brother subclade R1b-L23 was found in Eastern Yamnaya while R1b-L51 ''appeared somewhere in Anatolia''.

You really have a problem with the concept of time. and space, and geography. But you love Snps and statistics (what is that saying about torturing data?) that I grant you.

Those Balkans Farmers were in existence at least, at least 3,500 years before any Yamanaya or CWC even existed and Just 1000km from every theory, hypotheses or whatever any one is being talking here for the last 5 years. Yes, lol and behold even steppe theory! 3.500 years is freaking long time.

So, yeah, its a safe bet they had impacts on the yamanaya and CWC ancestors... and lets focus on seeing what their impacts were also and apart from steppe, on the late neolithic/chalcolithics in anatolia middle east, ararat, armenia, and so forth that suddenly show EHG (and I am not making any assumptions).

Maybe EHG in most of eastern europe (outside Russia) were I and not R1! That could explain why eastern europe and scandinavia still have a lot of I while western europe doesn't. And also would explain why the balkans have low ammount R1 and high ammount EHG admixture. Think about Serbia, very low R1, plenty of EHG, did the farmers have yamna wifes? Or EHG was there without R1 y-dna?

Yes, I think this has consequences mostly for the Balkans. Having significant admixture from SHG/EHG-like populations during the Neolithic does change whatever model one would choose for these populations.

How they might have contributed to other Europeans, if at all, is harder to say without even seeing the details of these samples.

It'll be interesting to see about Y-DNA. These foragers had to be related to whichever ones were in Ukraine. So whether they are R1a/b or not might be relevant too.

I don't think an "All EHG from the Neolithic Balkans" will be viable at all or "Most EHG", but it seems like it could perturb the exact %s over a model of just Iberia_MN / Ballynahatty + Yamnaya alone.

Not to a very large degree. For an example, just, say Yamnaya was 60:40 EHG:CHG, then Chalcolithic Balkans 20:80 EHG:AN (and an extreme case), then Chalcolithic West European farmers at 20:80 WHG:AN.

Then if you're estimating Yamnaya contribution to LNBA in West Europe, and they work out like 30 EHG, then that's too much for Balkans alone, but including Balkans might allow a slightly better fit than a 50% Yamnaya, if that doesn't quite fit for CHG.

Maybe EHG in most of eastern europe (outside Russia) were I and not R1! That could explain why eastern europe and scandinavia still have a lot of I while western europe doesn't.

The I2 in Eastern Europe is from WHG.

@Matt

I don't think an "All EHG from the Neolithic Balkans" will be viable at all or "Most EHG", but it seems like it could perturb the exact %s over a model of just Iberia_MN / Ballynahatty + Yamnaya alone.

How is this possible unless the Corded Ware and Yamnaya groups that moved west detoured through the Balkans and picked up a lot of admixture from the descendants of the Balkan farmers that still had some of the EHG/SHG-like Balkan forager admixture?

I know that some Yamnaya groups moved into and through present-day Bulgaria, but the Corded Ware expansion went straight into Central Europe from Eastern Europe.

Yeah, seems unlikely to matter for Corded Ware (at all) on the geographic basis. What about Bell Beaker and Bronze Age Hungarians tho.? Those are groups that may be ancestral to European LNBA and present day as well? Impact on model fit would be very small though.

Not sure if this was directed at my comment about path of least resistance but for a large n they don't need to figure it out in advance. Even if a population tries to expand in multiple random directions the ones who randomly pick the path of least resistance will succeed more often so probability (mostly) produces the same result anyway.

Yeah, the ancestors of the EHG-like Balkan foragers may have moved to the Balkans from north of the Black Sea, and some of their admixture maybe hung around the western Black Sea area into the Bronze Age. So they may have contributed something to western Yamnaya and Corded Ware.

PgmNr 1038: Genetic adaptation and Neandertal admixture shaped the immune system of modern human populationsWe find that marked differences in immune responses exist within and between human populations, due to the contribution of multiple cis- and trans-acting regulatory variants. These include, for example, a TLR1 trans-regulatory hotspot associated with the expression of a large gene network of 101 genes, which decreases the expression of multiple pro-inflammatory genes in Europeans only. (..) Specifically, our analysis uncover attenuated Inflammation as an evolutionarily advantageous mechanism, as attested by the cases of TLR1, CCR1 and DUSP14. Finally, we demonstrate that admixture with Neandertals has shaped monocyte gene expression by introducing regulatory variants into European genomes that affect preferentially responses to viral challenges, and report novel cases of adaptive introgression, as highlighted by the cis-eQTL regulating PNMA1 in response to influenza infection.

PgmNr 1046: An introgressed Neanderthal haplotype at the alpha-defensin CNV.We identified a putative Neanderthal introgressed haplotype in present-day European populations. Using a molecular clock approach, we found that the upper limit of divergence time between the putative introgressed haplotype and Altai Neanderthal haplotype (220,000 ybp)was more recent than the lower limit of the divergence between other human haplotypes and Neanderthal (670,000 ybp). (..) A global distribution of the introgressed haplotype showed that it was mainly distributed in Europe and Indo-European related populations, but absent from African and East Asian populations. We therefore believe this pattern to be a result of a European-specific introgression, after the divergence of Europeans and Asians. The frequency of the introgressed haplotype in European populations is 17.7%, with a higher frequency towards Western Europe. We further evaluated natural selection in this region, and found evidence of positive selection from the integrated Haplotype Score (iHS).

PgmNr 1071: The borrowed fitness: Adaptation of modern Eurasians through archaic hominin introgressionIn the Eurasian samples of 1000 Genomes Project, the contribution of Neanderthal introgressions to modern human was estimated at about 1% across the genome. High frequency (>30%) of Neanderthal alleles was found in genes related to inflammatory response, epigenetic modification, spermatogenesis, neuron development, etc. In particular, Neanderthal introgression in HYAL2 and MC1R, two genes related to response to sunlight, may have helped adaptation of East Asians in local sunlight. In addition to Eurasians, we found that 78 Denisovan introgressions in Papuans and aboriginal Australians were under positive selection, involving genes related to evolutionarily important functions, such as fertilization, cold acclimation, brain development, immunity, etc. Denisovan introgressions also introduced at least 121 missense alleles into Papuans, which are also located in evolutionarily important genes involved in female pregnancy, visual and smell perception, wound healing, and metabolism. These findings suggested that introgressions of archaic hominins may have substantially facilitated adaptions of modern humans to the new environments.

Are you talking with me? I was saying that EHG outside Russia had I2, EHG is WHG and ANE. Sorry, where is the contradiction?David, what is your explanation for the low amount of R in the balkans and in Scandinavia? (compared to their high ANE admixture) Why Serbia and Scandinavia have so much I whereas western europe doesn't?All I'm saying that we can't exclude the existence of populations that were mostly WHG/EHG (with some EEF) and with no or little R1 that lived in eastern europe and scandinavia right before IE started moving outside Russia. The only way to prove it is to find EHG/WHG mix in the bronze age that didn't have CHG or high amount of basal (and didn't have high percentages of R). That will explain the low amount of R but the high amount of EHG in scandinavia, the same goes for Serbia, and other places with plenty of ANE and very low R (the balkans). I mean, in Serbia ANE% is proably three times the R%. It's like the opposite of Spain, but nobody is asking why.

"If Neolithic Farmers from around Bulgaria (that seems to be from where the samples might be from) had up to 30% SHG/EHG admixture, that would have significant consequences in our understanding of LN/BA populations."

My guess is after the megafauna died out the mammoth steppe dudes drifted south. In the West (SHG?) they drifted to the coast and mixed / fought with the WHG while in the East (EHG?) the Black / Caspian Seas provided a livable niche that kept them further north.

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Azarov Dmitry said...

"This obsession with steppe homeland is so ridiculous. It’s based absolutely on nothing."

Ultimately it's based on the military advantage of horses at a crucial juncture in history when the farmers were still low density - which boils down further imo to the real crux which is the military advantage of having a population who spend ten hours a day in the saddle.

(which at the same time implies Scottish / Irish kilt wearing is a clue to something)

All I'm saying that we can't exclude the existence of populations that were mostly WHG/EHG (with some EEF) and with no or little R1 that lived in eastern europe and scandinavia right before IE started moving outside Russia.

Well, I reckon your jaw will hit the floor when you see what was in Eastern Europe west of Russia just prior to the Indo-European expansion. But there's no point continuing this discussion now.

David, what is your explanation for the low amount of R in the balkans and in Scandinavia? (compared to their high ANE admixture) Why Serbia and Scandinavia have so much I whereas western europe doesn't?

Y-haplogroup frequencies won't always correlate perfectly with levels of ancient components, because Y-haplogroup frequencies are more likely to be affected by recent founder effects and drift.

However, some Scandinavians probably have inflated ANE from Finnish and even Saami admixture. Serbs, on the other hand, probably get extra ANE from extra CHG which arrived in the Balkans independently of steppe movements, with people like the Anatolia Chalcolithic sample, and was probably initially associated with Y-haplogroup J2.

"So in that case who will have the advantage? farmers or pastoralists?. Farmers generally are more thick in nos."

Yes, hence a dramatic pastoralist population turnover would have to occur at the time where either- the farmers were widespread but hadn't yet developed high population densityor- after some kind of population crash possibly because they hadn't yet figured out how to preserve soil fertility away from land that did it naturally (i.e. self-draining loam soils replenished naturally each year by river flooding).

PgmNr 1156: Evolutionary history of Tibetans inferred from whole-genome sequencing. We demonstrate that Tibetans and Han Chinese likely first diverged in the middle Paleolithic period between 44 and 58 thousand years ago, but with substantial gene flow until between 3 and 11 thousand years ago; both populations have undergone large expansions since they diverged. Using a recently developed statistical approach to detect archaic admixture, we show that the Han and Tibetan genomes each contain about 0.4% admixture from ancient Denisovans. (..) Our haplotype analysis found that the advantageous Tibetan EPAS1 haplotype contained a large number of variants shared with the Denisovan genome, consistent with previous reports, but with an ancient gene tree divergence between the Tibetan and Denisovan EPAS1 haplotypes of about one million years ago. With the exception of EPAS1, we observed no evidence of positive selection on Denisovan-like haplotypes, strongly suggesting that adaptive introgression from a Denisovan-like ancestor was a unique occurrence in the evolutionary history of Tibetans.

PgmNr 1088: Ancestral origins and genetic history of Tibetan highlanders.Analyses of deep sequenced genomes (30–60×) of 38 Tibetan highlanders and 39 Han Chinese lowlanders together with available archaic and modern human data allows us, for the first time, to comprehensively characterize the ancestral makeup of Tibetans and uncover their origins. Tibetans arise from a mixture of multiple ancestral gene pools but much more complicated and ancient than previously suspected. They share the largest proportion of genetic ancestry with modern human populations from East Asia, and to a lesser extent with those from Central Asia, Siberia, South Asia, western Eurasia and Oceania. Moreover, non-modern human sequences, possible inherited from Neanderthal, Denisovan and other unknown ancient people, comprise ~6% of the Tibetan gene pool and elevated in some genomic regions. In particular, highly differentiated sequences harbored in the highlanders’ genomes were likely inherited from multiple non-modern human ancestral origins, and maintained in high frequency by natural selection.

PgmNr 1083: Identifying adaptive variants and tracing their ancestral origins in Tibetan highlanders by whole-genome deep sequencing.We build a list of putative adaptive genetic variants (AGVs) including 48 missense and 7 loss-of-function variants which affect 29 genes related to hypoxia adaptation. The majority of the AGVs are significantly correlated with altitude. Notably, we observe two highly differentiated genomic regions which show their elevated archaic ancestry and typical anatomically modern human origin, respectively. In both genomic regions, the sequences harboring AGVs in Tibetan highlanders trace their most recent common ancestor (MRCA) back to the time predating the Last Glacial Maximum. Our results indicated a much more complicated mechanism of human adaptation to altitude than previously thought, with key adaptions to the extreme environment contributed by pre-LGM populations.

PgmNr 1100: Genetic structure in the Sherpa and neighboring Nepalese populations.We identified clear substructure across populations resident in the Himalayan arc. The genetic structure broadly mirrors the geographical features of the region. Ethnic subgroups within Nepal show distinct genetic structure. We detected differential proportions of ancestry from northern Himalayan populations across Nepalese subgroups, with the Nepalese Rai, Magar and Tamang carrying the greatest proportions of Tibetan ancestry. The Sherpa appear as a remarkably isolated population, with little gene flow from surrounding Nepalese populations. We confirmed the presence of an ancestral component that appears specific to high altitude populations of the Himalayas and is enriched in the Nepalese Sherpa.

PgmNr 1057: An investigation of the evolution and worldwide distribution of the READ1 element in DCDC2 and its contribution to language development.READ1, located in intron 2 of DCDC2, is a multi-unit transcriptional control element with approximately 40 known alleles. Previous research has shown that repeat unit (RU) length groupings of alleles of READ1 (RU1-1 and RU2-long) are associated with RD [reading disability]. (..) We also examined the worldwide distribution of READ1 alleles in 58 populations among five different continental groups and discovered correlations between the allele frequencies of READ1 and several attributes of the languages used in a given culture: phoneme number (RU1-1, p = .006), consonants (RU1-1, p = .0151 and RU2-short, p = .0179), tones (RU1-1, p =.0084), and ratio of consonants to vowels (RU1-1, p = .0091; RU2-long, p = .0056; rs9461045T, p = .0104). We also showed a significant interaction between the SNP rs9461045 (a SNP in the KIAA0319 risk haplotype) and READ1 alleles in this correlation. The higher the proportion of RU2-long in the population, the stronger the negative relationship between the derived allele T of SNP rs9461045 and the number of consonants in a population. We hypothesize that alleles of READ1 may have influenced the type of written and spoken language developed by a given culture.

PgmNr 208: Global shared natural selection for increased stature in recent human history. Applied to data of ~3,000 genomes from the UK10K project, we identified signals of polygenic adaptation during the past ~2,000 years in ancestors of modern British that affected several traits. Above all traits, we found a strikingly strong signal for increased stature.Here we extend this work in two dimensions. First, we use SDS to analyze whole-genome-sequencing data for additional populations. In particular, we examine all 15 non-admixed populations from the 1000-genomes project (including populations from Africa, East Asia and Europe). Compared to the large UK10K cohort, SDS applied to the smaller samples of 1000-genomes (~100 individuals) detects adaptation on a larger timescale, of approximately 7,000 years. Second, we extend our method to infer the temporal dynamics of polygenic adaptation. We focus our analysis on the signal for polygenic adaptation of height.

PgmNr 1064: Testing differential genetic flow across Sinai and Bab-el-Mandeb.It has been established that SNP evidence suggests that the dominant out-of-Africa movement occurred across the Sinai rather than Bab-el-Mandeb, even though recombination event data suggests stronger admixture across Bab-el-Mandeb. (..)Our data supports the conclusion that the dominant genetic flow occurred over Sinai, with subsequent admixture primarily by that route.

PgmNr 1177: Fine-scale population structure in southern Africa reflects ecoegeographic boundaries.Our analyses reveal fine-scale population structure in and around the Kalahari Desert. Surprisingly, this structure does not always correspond to linguistic or subsistence categories as previously suggested, but rather reflects the role of geographic barriers and the ecology of the greater Kalahari Basin. Regardless of subsistence strategy, the indigenous Khoe-speaking Nama pastoralists and the N|u-speaking ≠Khomani (formerly hunter-gatherers) share recent ancestry with other Khoe-speaking forager populations that form a rim around the Kalahari Desert. We reconstruct earlier migration patterns and estimate that the southern Kalahari populations were among the last to experience gene flow from Bantu-speakers, approximately 14 generations ago.

Here is the thing. Bulgaria in 4200 BC shows up to 30% EHG-like admixture, that's quite south, and the region didn't even have any Mesolithic sites (apart from Bulgarian tip of Dobrudja). So imagine what the C-T culture would look like- 1000 years later, & from a region with abundant Mesolithic habitation ?

@Grey: "fascinating". Indeed. Though I wonder whether we have a one-directional causality here (genes shaping language), or a feedback loop, i.e. linguistic capability in certain cultural contexts being positively selected, thereby enhancing the occurence of specific READ1 variants. Anyway, the relation of genes and language deserves further in-depth investigation.

Coming back to my earlier postings on Neandertal and Denisovan introgressions, I wonder whether, similar to uniparental markers, we may use such introgressions to trace population movements. Alpha-defensive CNV, restricted to (Indo-)Europeans, e.g., should allow to discriminate between basal and non-basal Eurasians. Those 78 positively selected and 121 missense Denisovan alleles in Papuans might help to trace Out of Papua geneflow (Andamans? Americas? Madagascar/ E. Africa?); etc.

Davidski: Bell Beakers and the Bronze Age Hungarians don't get their eastern shift from Balkan farmer admixture. They get it from the steppe.

You seem really confident that there's no significant Balkan farmer admixture in BB or Bronze Age Hungarians, as well as the steppe admixture we know to be the case. Not sure why you're so sure but not going to argue the case here further (since we don't know much about the paper anyway!).

One thing that is interesting in the abstract that we haven't talked about is "We show that many of the adaptations related to diet and immunity that later become common in Europe were already present in early Balkan farmer populations, but not at high frequency. " so I wonder if this is a reference to LP (though there are other diet adaptations - FADS1)

In agreement with earlier studies, we find that selection for increased height has recently elevated in Northern Europe. However, surprisingly, we find that all examined populations, independently across continents, had experienced detectible selection for taller stature. This apparent global shared selective pressure could not have been detected with earlier comparative approaches. Our work thus sheds new light on recent evolution of human complex traits.

That's very interesting in light of the selection results in Sardinia for shorter stature (e.g. http://www.ncbi.nlm.nih.gov/pubmed/26366551), and suggestions of the same in early Iberian farmers (and Great Andamanese). Clearly cross checking against the large Sardinian whole genome reference is called for.

So I wonder if there is some degree of balancing selection here going on, where the general pattern is selection for taller stature in world populations, while in some populations there is a countervailing selection for alleles which are stature reducing but have some health benefit?

It's also unusual in that bioanthropometry doesn't indicate modern people are necessarily much taller than in the UP or Mesolithic. So I wonder if it is the case that many height alleles they find are actually "Height if you're fed on an agricultural diet" alleles (which would fit with global selection after transition to agriculture).

You seem really confident that there's no significant Balkan farmer admixture in BB or Bronze Age Hungarians, as well as the steppe admixture we know to be the case. Not sure why you're so sure.

Because we have a number of Hungarian and even some Greek samples now from across space and time, and none of them shows anything like that, except for Hungary_HG's somewhat inflated allele sharing with Motala_HG.

If the Bell Beakers and Bronze Age Hungarians do have Balkan farmer ancestry, then they got it from their steppe ancestors. This is indeed possible, but if so, it won't be a lot.

I have another theory about the origins of the Bronze Age Hungarians. I think some of their ancestors arrived from some part of northeastern Europe with very high levels of WHG, which was Indo-Europeanized from the steppe during the Late Neolithic.

@Matt: "bioanthropometry doesn't indicate modern people are necessarily much taller than in the UP or Mesolithic. So I wonder if it is the case that many height alleles they find are actually "Height if you're fed on an agricultural diet" alleles (which would fit with global selection after transition to agriculture)."

Not so long ago, there was an article on biovrix that pointed at a possible connection between yDNA and (male) stature showing up in British panels. Both R1 and I seem to be associated with taller stature, R1 also with higher BMI. Findings weren't statistically significant, though, for an insufficient number of samples other than I and R1.If true, the late CA/ EBA star-like expansions of R1 and I1 would have helped to bring Europeans back to UP/ Mesolithic statures, reversing the impact of smaller EEF, possibly related to their yDNA. Taller stature is more of an advantage for hunters (and, later in history, warriors) than for farmers. The latter will better survive periods of malnutrition (harvest failure) if they are smaller and have the ability to accumulate fat reserves instead of quickly mobilising energy for fighting/ running. Substantial height differentials are well documented between African hunter-pastoralists (Nilo-Saharans, Bantu, Tutsi, etc.) and farmers (Ewe/Fon, Igbo, Hutu etc.).However, the picture is most likely more complex. Availability of amino-acids, be it from dairying or stronger intake of meat/fish, clearly plays a role, and, as the linked abstract indicates, there seems to be positive selection on height independent of the influence of certain yDNA hgs.

MattIts interesting, because the Robust Mesolithic European foragers were coming from a markedly diminished background. I mean, compared to pre-LGM stature, the foragers in both east and Western Europe immediately post ice age were smaller- as perhaps some kind of adaption : small limbs lose less heat. They must have beefed up again by the Late Pleostocene

@Dave: "If the Bell Beakers and Bronze Age Hungarians do have Balkan farmer ancestry, then they got it from their steppe ancestors". Your K10 shows Hungary_EBA (I1502) sample with little Steppe admix (3.3% CHG), though quite some EHG (16.6%) that apparently entered from a different direction, possibly relating to SHG.

"I think some of their ancestors arrived from some part of northeastern Europe with very high levels of WHG."Steppe admix increases towards Hungary-LBA (I1504), which has 11.2% CHG. That 8% CHG gain against EBA corresponds to a 7.5% decrease in WHG, which makes an arrival from NE Europe quite unlikely. Rather, I think we are dealing here with CHG influx from the Mediterranean during the MBA/LBA.

Current Hungarians range at 27.7% CHG/ 13.9% EHG, which is +15.5% CHG, -2.3% EHG compared to Hung_LBA. So, the main Steppe influx, strongly CHG-loaded, seems to have occured after the LBA. Skyths, Avars, Magyars come a/o to mind here. Especially the Avars, originating from the North Caucasus, should have been heavily CHG-loaded.

Interesting is a look at Hung_CA (I1497). It is from a Baden Culture context, ca. 2800 BC, as such without cultural affiliation to Steppe/ CW, and predating Hungarian BBs. This sample shows up in Dave's K10 as 3.3% CHG (as Hung_LBA), yet only 1.6% EHG (14.5% less than Hung_LBA), and 13.8% WHG (17.1% less than Hung_LBA)

This indicates:(a) CHG presence, at a low yet measurable level, in Central Europe already prior to Yamnaya;(b) Substantial population replacement in Hungary between 2800 and 2100 BC. When assuming that the incoming populationn contained no EEF ancestry, we are talking about 40% replacement. If the newcomers held some EEF ancestry, the replacement rate goes up further (53.5% at 20% EEF ancestry w. the newcomers).(c) The non-EEF part of the incoming population was something like 58% WHG, 38% EHG, 4% CHG. This doesn't have much to do with Steppe, though a little bit of it may have been included. Instead, it looks like slightly western-shifted SHG. My best guess would be non-CW, GAC-related populations of the CE plains, e.g. from Kujawya, Elbe-Havel, Lower Middle Elbe or the Netherlands. Unfortunately, we still lack aDNA from any of those.

"Here, use the Basal-rich K7 spreadsheet and nMonte to calculate this sort of stuff...It isn't that informative here, as it compresses four of the K10 dimensions (WHG, EHG, EEF, CHG) into three (ANE, basal-rich, Villabruna), and, more specifically, makes it virtually impossible to distiguish EEF and CHG.Nevertheless, at a first glance, it shows something like

Essentially the same story: ANE and Villabruna go up strongly between CA (Baden) and EBA, which can't be explained by Steppe influence. The LBA has undergone influx of a slightly ANE-enhanced basal-rich population, which also can't be predominantly Steppe, but instead looks mostly Mediterranean (Cypriot-like). Current Hungarians are shaped by later Steppe influx that is more basal-rich (CHG) than Yamnaya.

@Rob: We are talking about some 40-75% population replacement between Baden (ca. 2,800 BC) and the EBA (ca. 2,100 BC), for locations that lie 50 km apart from each other in NC Hungary. Can't be pre-Baden, nor, for the size of the replacement, just exogamy. If exogamy with Yamnaya played a role, those Danubians furthermore must have refuted all the CHG females and just gone for the EHG ones.

The Hung_EBA sample stems from a Makó Culture context. Unfortunately, Makó Culture seems to be one of those "Transcarpathian Phantom Cultures" that show up in all chronologies without much further substance/ research associated to it. You may, e.g., try to check the "Oxford Handbook of the European Bronze Age" for details on the Mako Culture - I couldn't find anything specific, but may have overlooked a note here and there. While they left some ceramics (https://commons.wikimedia.org/wiki/Category:Mak%C3%B3_culture_in_BHM_(Buda_Castle) ), there seem to be hardly any settlement traces, and only and handful of burials.

"Is there any evidene for such a movement ?"Well, the Hungarian_EBA aDNA..

Some authors regard Hungarian BB as resulting from migration up the Rhine and then down the Danube.There is a late 3rd mBC cultural trajectory from the Mierzanowice Culture (L. Poland) via the Nitra Culture (Slovakia) into Northern Hungary (Hatvan Culture) that constitutes a clear break to preceding CW / Epi-Baden.http://www.geochronometria.pl/pdf/geo_19/geo_19_06.pdf

Plus, Hungarian EBA has cultural pointers to the Harz, such as "Totenhütten" (post-built inhumation houses) and certain tin crucibles. However, those pointers may relate to later periods (Unetice).https://de.wikipedia.org/wiki/Totenh%C3%BCtte_von_Benzingerode

The closest proxies for the admixing population in Hung_EBA that I could identify were CHG-poor BB and Unetice samples from the Elbe-Saale Region, e.g. I0803 (Eulau - Unetice). However, while relatively EEF-poor, they still come in with way too much CHG/ EHG. A mix of EEF-poor German BB/Unetice, and a Steppe-free population, e.g. Iberia_CA, might do the trick, though.

@Nirjhar007I think Balkan Hg's will show R1b ( extinct branch of course ) , don't think will be any R1a but doesn't matter its too old ;) .

I’m sure we’ll see a lot of R1a-YP1272 folks that came in Balkans (Chernavoda, Baden and Ezero cultures) from Pontic steppe (Sredny Stog culture) and after 3300 BC some R1a-YP1051 folks migrating from Anatolia (Ezero culture). R1a-M417 guys came later (probably only after 2600 BC or later). And of no doubts we’ll see some R1b-L51(xL151) folks coming from Anatolia.

@AzarovD, OM: Sorry to disappoint you both a bit:The earliest L23 found so far comes from Yamnaya Samara - I0429, ca. 3,100 BC, and it is already ZZ103, i.e. a L51 sister clade. Now, one would need a pretty convincing story - archeologically - to link Samara to Anatolia.The earliest L51 is the famous Quedlinburg Bell Beaker (I0806, ca. 2250 BC), the closest relative to the Irish Rathlin I (ca. 1950 BC). BB is still Chalcolithic, not Bronze Age.We have a bit of Baden/ post-Baden aDNA already. No R1a so far (and little indication aDNA-wise of enhanced Steppe contact), but one R1b from Lánycsók, HU (ca. 2700 BC). Lánycsók lies next to the western bank of the Danube, near the Croatian border, and is much closer to Villabruna than to Samara.

I took the Balanovsky abstract linked by Dave as an opportunity to check on the ocurrence of R1b1a2a1 (L51) vs. R1b1a2a2 (ZZ103):

The geographical split between CA/Scandinavia and Steppe/Caucasus becomes apparent. Of course, there is quite some aDNA that has only been determined to R1b1(a), and new samples may change the picture. But for the moment, it seems as difficult to link Yamnaya R1b-ZZ103 to BB/ Battle Axe R1b-L51, as to predominantly R1a Corded Ware.

Ultimately, it all comes down to the question where R1b1 diversified. - Anatolia? Hasn't AFAIK yielded a single R1b1 aDNA so far;- Caucasus? There is the Kura-Araxes R1b1 from Armenia (I1635, ca. 2550 BC), but it was negative for M269, so it can't be ancestral to L51, nor to ZZ103.- Steppe? No aDNA evidence for L51, and very little L51 among current populations.

What does this leave us with:- Earliest R1b1 from Villanova, NE Italy- Earliest R1b-V88 from El Trocs, EN Iberia- R1 (xM269, xR1a1a) from Baalberge Culture, ca. 3550 BC- R1b1(xM269) in Kura-Araxes, Armenia- L51 predominating in Central Europe, ZZ103 in the SteppeTriangulation between these endpoints takes us to somewhere in the Northern Balkans or the Carpathian Basin. And, apparently, M269 (xL23) seems to peak in Kosovo,

Hopefully, the upcoming paper on neolithisation of the Balkans will provide some further insight.

But for the moment, it seems as difficult to link Yamnaya R1b-ZZ103 to BB/ Battle Axe R1b-L51, as to predominantly R1a Corded Ware.

One of the Battle-Axe remains belongs to R1a-M417.

And it's not difficult at all to link Corded Ware to Yamnaya, because the Corded Ware cluster extends from the Afanasievo/Yamnaya/Poltavka cluster into Europe, which is of course completely in line with formal stats.

Thanks for reply; I agree with your observations on the whoe, back to HungaryOn present balance, I'd favour the arrival of the distinct-signatured eastern shift in Hungary to miscegenation with western Yamnaya, who are probably more WHG shifted western version of Kalmykian types; and/ or Cernavoda, which also appears from the Danube -Dniester interfluve.

Lastly, we need to be careful about making conclusions from modern populations, esp drifted recently expanding ones like Kosovars which have high feuencies of EV13 ans Z2013 due to founder -effects. With regard to modern Hungarians, I suspect, for most part, from whatever pre-Magyar groups were resident in the Carpathian. Being Uralics from the Volga (ultimately) I doubt the Magyars had much CHG, even after the sojourn with the Khazars; nor would the Scythians et al left much, as they're basically "extinct" groups. The Balkans is very complex, and modern groups shouldn't be taken as proxies for Bronze Age ones

"Thanks for information... so much ado about nothing..."Someone said that when it was clear that the new subclade found from Balanovsky et al (GG400) is only Z2103, and it should be so, because the subclades present in Yamnaya and in Middle East, and less (not a little, seen that I belong to them too) in Western Europe, are subclades of Z2103.It isn't frequent to may ask one of the authors of the paper (in this case Vadim Urasin), even though in other blogs many illustrious geneticists usually write, even though hidding themselves beyond a nickname. Vadim, of course we'll read the paper with very much interest, but have you never read what I am writing also on this blog? I doubt that Z2103 was born in Eastern Europe, lacking all the upstream subclades there but present in Italy and Western Europe, I doubt above all that the Western European Z2103 subclades descend from Yamnaya, if not some minor subclade like CTS7763, but very likely not my Z2110 neither the huge CTS9219, being the upstream CTS7556 and Y5592 above all from West, thus they may be a back migration from West. Also about L584 and L277 I wrote a lot.Of course when YFull linked the Sardinian Z29823 not to Z2103 but to Z2110 and with the Sephardic (and Iberian!) CTS699 (what also smal accepted and I hope it will be in his next tree), I understood that you worked well.Hope that in the paper, with many other samples from Eastern Europe deeply tested, I may answer all the questions I raised.

@FrankNUltimately, it all comes down to the question where R1b1 diversified. - Anatolia? Hasn't AFAIK yielded a single R1b1 aDNA so far;- Caucasus? There is the Kura-Araxes R1b1 from Armenia (I1635, ca. 2550 BC), but it was negative for M269, so it can't be ancestral to L51, nor to ZZ103.- Steppe? No aDNA evidence for L51, and very little L51 among current populations.

Main splits of R1b subclades occurred in Caucasus, Anatolia and Corsica.

Dear Azarov, from your profile it seems that you live in Moscow. Your name is Russian, but your surname is russified. Are you perhaps a friend of Anatole Klyosov?Your map is ridiculous, because of your pathway of hg. R1b there isn't any witness either in the aDNA nor in to-day diffusion. The first R1b found in Iran has been 2800 years ago F38, an R-L584 come from South Russia with Medians, who were Indo-Europeans and had nothing to do with the people of Old Iran, of the Zagros mountains.

@DavidskiFrom yours Basal-rich K7 table I see that Villabruna component of Iran_Neolithic = 0% but Iran_Chalcolithic = 7-11%. This suggests that there was an influx EHG-like component into the Chalcolithic Iran.

@Davidsi.And, Respectfully, I do think you have a neurocognitve Impairment (ie, you are a ...).

the adna we have is a couple of dots in a future to clarify highly complex picture...the adna we have is highly biased to the most northern hemisfere (i.e. ICE/cold preserving) and is a factor to have in mind while infering.the adna we have is at its infancy compared to say 200 years of anthropology and might actually just represent that specifically individuals and not the all culture (we just saw that with BB germany).

The imparment part? - you just got kicked in the balls by EHG found in eastern Europe almost 4000 years prior to Yamnaya, 700 miles from Germany, 800 miles from the steppe. and your comment is "unless they contribute to Yamnaya".

The point I am trying to make is for you to tone down and not sound like a prick.

secondly that, like all sciences in its infancy, more often then not are wrong than right. Especially, if you replay is " main models", I work with models and let me tell you to put models in a certain dark place. Models represent the shit you feed into it and nothing else.

thirdly.just trying to figure out, how do you know that all the inference you scream/shout here about Yamnaya in western Europe gemone does not relay as well or better with the Balkans Farmers that were actually in europe 4000 years earlier and not to an no archaeological migration of Yamnaya?

fourthly - Nobody dispute the CWC influence in europe (mostly eastern and central) but how do you know that that EHG/putative Yamnaya in CWC does not actally derive from those Balkans Farmers being them in the region 4000 years longer that the posterior CWC/Yamnaya.

THe list of points could go on forever, move to anatolia, and middle east (that shows chalc EHG and WHG) and so forth...

Nobody dispute the CWC influence in europe (mostly eastern and central) but how do you know that that EHG/putative Yamnaya in CWC does not actally derive from those Balkans Farmers being them in the region 4000 years longer that the posterior CWC/Yamnaya.

That might be an issue in the la la land that you inhabit, but in the real world, it's really fucking simple.

Run a PCA with a lot of ancient samples, and look at the relationships between them and present-day samples.

Guess what? The Corded Ware cluster emerges out of the Afanasievo/Poltavka/Yamnaya cluster, and stretches into Central Europe, towards modern Europeans.

Then we look at the data more closely using formal stats, model-based tests, and whatever else that's available, including uniparental markers. And what do these show? They show that the PCA is correct.

Really not very difficult. In fact, the more ancient DNA that comes in, the more obvious these things look. You just wait for the Baltic paper sunshine.

@ Davidsi,Oh yeah? - And where are those 8000 years old Balkan farmers in that PCA plot?

When they "cluster" (and some of your PCAs to show CWC very well running away from that cluster) the Balkan Farmers with them, what will be your story? Yamnaya travel back in time?

Secondly, who predicted we would find Such a EHG in Anatolia next door? Balkan my ass. If its Bulgaria is "Anatolish"! Black sea shores makes it Next door. And how come CHG Kotias (next door Black sea shore) had such an amount of EHG?

Olympus, you're making out like those Balkan farmers have been introduced as some sort of a game changer. But they haven't.

We can reasonably figure out how they will cluster, simply by positing that with 0-30% SHG/EHG-related admixture they'll look like more northeastern Anatolian farmers.

How will that change the fact that Corded Ware cluster close to Yamnaya and look like over 70% of their ancestry is from Yamnaya? I don't see how unless some of those Balkan farmers are basically Yamnaya.

But of course they're not basically Yamnaya. I can tell you that Corded Ware colonized most of Eastern Europe from the steppe, and the people living today in most of Eastern Europe are largely of Corded Ware origin. You can take that to the bank.

"Ultimately, it all comes down to the question where R1b1 diversified. - Anatolia? Hasn't AFAIK yielded a single R1b1 aDNA so far..."

Which explains that the spread of agricutlure and the diversification and distributioon of R1a/R1b has nothing to do with the spread of agriculture.

Which seems rather contradictory to the present distribution of farming, diary and LP.

There's a row of factors ascribing to a relation between the spread of cattle-farming and the distributin of R1a/R1b.

Moreover there's a clear-cut dominance of mt-dna H showing up simulatniously. For some reason they both seem to spread along with a hightened ability to digest starch, lactose and milk-proteins. Still there's noone claiming mt-dna H to be "Anatolian".

So what g-e-n-e-t-i-c evidence is there to explain that "farming spead from Anatolia"?

If there's isn't any - why do reputed geneticians keep repeating this hypothesis as a mantra? Are none of them aware of that the archaological views on this subject are differentiated and still up for further analyzis and debate?

Stuffing empty dogmas can never be good science.

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On-side:

The gene respeonsible for producing amylase in the pancreas, to digest starch, is quite high in the "hunter-gather" from Loschbour (12) compared to the ENE from Stuttgart (14). Meanwhile the HG from Mottala came out at the bottom of the scale (4). Acccording to herrn Krause.

Which means Louscbour were clearly more adapted to terrestial veggies than Motala. Which still makes med wonder why Loschbour is a better proxy for "WHG" than Motala. Supposing that the terms in question have something to do with archaeology and common sense...

Regarding the initial branching of R1 into two separate dynasties - as R1a and R1b:

I think we have to look for a geographical zone where the climate as well as the natural resources needed for cattle-farming existed during the Mesolithic already. Then we may look for the areas that during ENE could succesfully feed larger herds of large cattles.

In that case we know that the occidental climate of the German and French plains - as well as the English lowlands - were very early. The oldest burial of an ox is known from the SW Baltics already 11.000 yrs ago.

Thus we should look between the Vistula and the Rhine to find the diversification of R1a and R1b.

Looking at the present diustribution it's nop doubt that we have a 'diversifying' are along the Vistula-Oder-area.

Which is why I have a problem with the plain assumption that R1 came out of anywhere but central Europe. Skilled anaklysts seem to agree, eventhough I'm not sure how sure they are about their own results...

@ Davidsy.No.it means that ALL the EHG you have been pushing as Yamnaya flooding into western Europe might actually not be the case of being ALL yamnaya hence reducing their legacy to something closer to what archeology shows (which is nothing if you cut CWC).It means that we have to recheck if CWC is actually 70% Yamnaya or just say 30% and the rest coming from some other much earlier source since we found closer to CWC one that actually was 30% EHG and over the course of 4000 years as we sample all those tens of north Caucasus cultures i that 4000 span we will probably find lots of EHG mixed with CHG that could be an added component to CWC later populations.

and why not, since we have a 8000 year old 30% EHG (that is 4000 earlier then Yamnaya/cwc) that was on the pathway of all out of Anatolia and Into Anatolia/Caucasus. Chalcolithic in those regions (and chalc there is 1000 years back Yamnaya/cwc) so lots of time to contribute to the mix of Europeans. who knows, we might just find other EHG rich that actually gave EHG/CHG to european populations. The landscape of DNA os full of holes that span large millennia.

Regarding CWC - No argue from me that CWC (WOMEN!) were part of the admixture that mixed in Bohemia with Bell Beakers and up Elbe river and ended up with heavy impact all over europe (wherever those dudes went). No Argue from me that CWC (MEN) had a huge impact on Unetice cultures and so may more that craft eastern europe dna landscape. And that CWC descendents got important exogamy with other cultures. Autosomally, it all mixed in southern europe later cultures and north Europeans Celtic (which was an insult)/ belgae and so forth.

@DavidskiHow will that change the fact that Corded Ware cluster close to Yamnaya and look like over 70% of their ancestry is from Yamnaya? I don't see how unless some of those Balkan farmers are basically Yamnaya.

But of course they're not basically Yamnaya. I can tell you that Corded Ware colonized most of Eastern Europe from the steppe, and the people living today in most of Eastern Europe are largely of Corded Ware origin. You can take that to the bank.

It’s a bit more complicated. First of all Corded Ware culture is a result of expansion of mostly western part of Yamnaya culture (Sredny Stog population (mostly R1a-YP1272) transformed as a result of cultural contacts with Maykop population (R1a-M417)). Secondly ancestors of the people living today in Eastern Europe came not from Yamnaya (first wave of migrants) but from Maykop -> Catacomb cultures (second wave of migrants).

Distribution of main R1a subclades in Eastern Europe by 3000 BC

http://s017.radikal.ru/i419/1609/b1/8ccb18c49f10.jpg

Distribution of main R1a subclades in Eastern Europe between 2900 – 2700 BC

http://s019.radikal.ru/i616/1609/aa/0af2b75656c3.jpg

Distribution of main R1a subclades in Eastern Europe between 2700 – 2200 BC

I've never said that It was. I only say that some folks (R1a-L664 and R1a-Z284) migrated from Maykop -> Catacomb cultures on the territory that was earlier colonized by R1a-YP1272 folks(migrants from western part of Yamnaya). That's all.

The Catacomb culture shows links ? moves to Middle Dnieper culture & Cracow-Sandomiercz group of CWC. In turn, the link of Catacomb culture to the Caucasus has always been discussed, but its always remained arguable.

Nor I suppose CHG is from the steppe.... but had a good portion of EHG. well that EHG sure travelled a lot, didnt it. Caucasus, balkans by 8000BC?... so lets leave the Yamnaya 5000 years later rest for a bit and see where else that EHG dude went in those 5000 years, shall we?.

Model after model have said Yamnya works as the eastern proxy who changed Europe after the Neolithic, we have no models saying the same about the Balkan Neolithic yet you whole heatidly think the Balkan Neolithic is a better proxy.

Are you time traveler who got bullied by Yamnaya children from the other side of the forest? I have no other expliniation for your magnetic dislike for Yamnaya.

Take real D-stats data for Motala_HG (SHG) and Karelia_HG (EHG), combine with real D-stats data for Barcin_N (Neolithic Anatolia) in proportions either HG:A 22:78 or 10:90. This produces the simulated populations.

Then I plotted these on PCA with the real D-stats that Davidski had produced.

This should work as a first approximation because combining D-stats data for Bichon (WHG) and Barcin_N reproduces the D-stats for the real Iberia Early Neolithic and Iberia Middle Neolithic populations, which should be analogous populations to what is under discussion except with WHG admix rather than SHG / EHG.

Alternately, you could do the same thing with Davidski's PCAs as well, if you wanted to.

The evidence thus far gives me the strong impression that r1b and r1b arose somewhere between central europe and siberia, and possibly connected to ANE.

As for you talking about Villabruna... You are talking about WHG (+ ANE?) group which is also shared by yamnaya but not the middle east. The thing you have to explain is how the yamnaya became L23 dominant when the variant arose only 1000 years prior.

"What bell beakers were 50% yamnaya?Not the "half breds" in Germany that we know were found 100 miles from the Bohemia BB group that we know were the sons/grandsons of CWC women, right?"

You arent going to get a large half breed population from a short time of mating with ONLY women. I dont care what label (bell beaker or CW) you give for the men that came from the steppe and entered western europe, but that is exactly what they did.

You sound exactly like the people i used to argue with who thought only r1a was eastern. Bohemian bell beakers were not the result of western bell beaker men. Desideri already showed us this.

If you are clever enough to ask me how I think the yamnaya became 50% caucasus like without (most likely) absorbing caucus like ydna i would argue that the time period of mixing was much larger and the yamnaya likely had a population boom after the mixture took place.

@Colin,So we are back to "might" and "likely"... pretty good. This is what Desideri, after 15 year publishing BB paper and 10 on Nm dental traits, is saying in her own words:

“the emergence of the Bell Beaker culture in the southern sphere resulted from the displacement of individuals from the Iberian Peninsula into Europe.The biological impact was recorded to at least Switzerland, and possibly also to Hungary. Thus, the Bell Beakers – small groups of individuals equipped with their material culture and know-how – formed the basis for Bell Beaker diffusion in this region ofthe phenomenon. - the situation in the eastern sphere is more complex.”

Then,

“The southern Bell Beakers are quite similar and the Swisspopulations can be strongly linked to their morphology. They form a highly uniform group. Theeastern Bell Beakers show a certain cohesion that seems, however, to have been less isolated”

@colin,They were half breeds. They showed Iberian Chalcolithic (everybody likes to forget) and showed overloaded CWC/eastern linkage. Of course CWC males / BB women were part of the exogamy that made the Eastern Group. That Eastern BB group were later part of Unetice and other (davidsky, davidsky, listen carefully) Bronze age cultures in the balkans... sure.And overall northern europe...sure. CWC guys rocked. Its just the worthless Yamnaya/Apple Iphone that get on my nerves.

If a group of 50 BB man steal 50 CWC women how do you think their Dna look like when they show up in a settlement 100 miles away?

What we need is someone to do the same as Desideri did in central/western france, Netherlands, etc... that would give a clear picture.

@colin,""If you are clever enough to ask me how I think the yamnaya became 50% caucasus like without (most likely) absorbing caucus like ydna i would argue that the time period of mixing was much larger and the yamnaya likely had a population boom after the mixture took place. """

Well wont ask, but I can tell you. Samara Yamnaya might (since we are allowed) have been part of the exodus of "my shulaveri" and those, living in Kotias land, had EHG and CHG. by 4,900 bc here in Kuban river and by 4000bc could be in samara river already admix with local loaded up EHG populations.why do you think Abashevskaya culture (the descendents of Samara Yamnaya) non metric skeleton and crania (dont trust, dont trust, ;) ) is a perfect match to the Kura araxes isolated in the mountains (so leftovers of Shulaveri)...

"If a group of 50 BB man steal 50 CWC women how do you think their Dna look like when they show up in a settlement 100 miles away?"

Lets do the math. The women were at most 3/4 yamnaya like and according to your theory the men had none. Their first generation is now only 3/8. Those offspring move 100 miles to the west and mix with a zero yamnaya like person, so that the second generation is 3/16 yamnaya like. Do you see the problem?

We have no method which can seperate Iberia Chalcolithic from other Middle Neolithic/Chalcolithic Europeans. Iberia Chalcolithic works as well as Irish Neolithic or Swedish Neolithic as an ancestor proxy for Bell Beaker Germany. There's nothing strongly supporting or not supporting Iberia Chalcolithic ancestry in Bell Beaker Germany.

There's also no convincing evidence Bell Beaker was part or not part Corded Ware.

It seems there is no link between South Caucasus and north until after Majkop & K-A began . In other words, the coastal strip of Azerbaijan was colonised by KA groups from the south & Majkop groups from the north, which must have been long separated cultures, filling in a sparsely populated "no man's land".

Can you explain to me which is the scientific principle as to Tuscans of to-day (after 4000 and more years) derive from Yamnaya of 4000 and more years ago and not the other way around, also after that I think having demonstrated that R-L23 is more likely born in Tuscany/Italy from R1b1-L389*, R-L297* (close cousin of Villabruna of 14000 years ago, found in Italy and not in Yamnaya?=

The way forward when it comes to unravellingMore recent admixture in related Pops is by establishing a good y DNA database. Formal stats reach their limitations here, unless we have exceptional quality data

"You would need to have Sardinians living in Tuscany today for this not to be true".

But one of the strong points of my theory of an "Italian" origin of hg. R1b1-L389+ and subclades (amongst many other Y-s and mt-), are just the data of the 1204 Sardinians tested from my friend Paolo Francalacci and his team of the Sassari University.Sardinia has, from Continental Italy and from Mesolitic,: R-V88 (M18 and V35: the oldest of the world), and also the oldest subclades of M269*, L23 (amongst them my Z2110, now a SNP discovered from YFull and studied by me and accepted from Sergey Malyshev (smal) in his last tree of to-day, links two Sardinian samples with the CTS699 subclade, found in Iberians and Jews: Jews think that Iberians are Crypto-Jews, I think that Jews are introgressed in Europe: 70%, 45% Italian), L51*, U152 etc., of course with old haplogroups pushed Westward like I-M26 and G-L91 etc etc.

Just forget Y-DNA? But my theory is just about Y-DNA and hg. R1b1-L23 in this case, found at Samara but without its upstream and downstream subclades! It will be easy to understand where Samara R-L23 did come from, just testing samples older than 5300 years ago and see if there are upstream subclades, what lack in all Asia and Eastern Europe.Also with my rough exams I have said that some hg. J older than Satsurblia will be found in Italy. This is science: to make previsions and experiments that prove or disprove that.Please, explain me what your data should demonstrate, in words for a newby, as I am in this matter.

dartraighe wrote:GioielloHow can EHG be a mixture of ANE+WHG?!!!!!! WHG is one of the regional variants and autosomal dna is region specific. That is why we all look a little bit different. The revenge of the HGs in western Europe is a myth.

"I am saying that WHG, ANE and EHG are three separate components. Villabruna had 15% ANE, as well as having mostly WHG, which shows that the Italian people had the ANE component in western Europe 14,000 years ago".

@Samuel Andrews Oh yes there is. Because IBerian Chalcolithic had as whiff of SSA dna that you dont find anywhere else. And BB, even the Freaking half Breeds in Germany had that Whiff of SSA. That was the inference Ryuenko and Davidsky danced to a couple months back.

@Colin,"anyways, i see no point in arguing on the basis of dental traits"

Right. Is like saying, "your honor I refuse to discuss the matter based on Footage because is not HDTV...!"

Meanwhile it looks like you are more then happy arguing on Models?! on statics inference of models?! -- Man, if you don't know a model represents nothing more than the model itself...! Its the Logic of Shit-in = Shit-out.

"It is a fact. EHG are somewhere around 40-50% WHG. But all of the WHG samples that we have are also part ANE, probably usually around 15%. This might just be a feature of the WHG clade because Villabruna has it too. So EHG are probably something like 75% ANE and 25% proto-WHG".About that "proto" we could discuss...

@Rob """"It seems there is no link between South Caucasus and north until after Majkop & K-A began . In other words, the coastal strip of Azerbaijan was colonised by KA groups from the south & Majkop groups from the north, which must have been long separated cultures, filling in a sparsely populated "no man's land"."""

Man, what I find most amazing here at Eurogenes is that so many have a problem with several concepts (time, space) but most, most of all with the concept of ... HUMANS.Rob, At a very narrow bandwidth of time (5000-4.800 BC) you find the vanishing of a whole culture. from Arukhlo to Mentesh Tepe or even a far as Aratashen, you have something like tens and tens of thousands of people. Clearly replaced by a diferent culture.

So, when do the settlements of the Kuban river, near the eastern shores of black sea start to popup as Svobodnoe and the all Zakubanye culture?

And what did those guys had? that the steppe didn't? Agriculture highly mixed with Pastoral and transhumance life style, right? The fauna is the same as south caucasus, and those settlements do have a lot of Shulaveri traits.

So if at same time you have a large number of population Vanishing 200 miles away from a large number of people appearing, especially having similitudes.. what is that in this Play station 4 world we here game is really suppose to mean?

It is possible, but the pre- Majkop, Sovodbojna phase looks like a gradual continuition of local Neolithic, still not extensively explore The big change occurs c.3800 BV, this 1000 years after your event

MALES - What Desideri shows it that bell beakers in the Bohemia actually showed males from the BB package (IE IBERIA males traits) and also males from CWC that do appear int the Bell Beaker package. So half breeds. Otherwise it made no point on making the distinction of CWC males and BB males, which she clearly does. It goes on to say that those BB MALES traits do not show at all in the Unetice Males, but CWC nm dental traits do indeed show up in Unetice.

FEMALES- What Desideri shows is that Both BB females and CWC Females showed in both cultures indication lots of exogamy and a strange fact that not even the BB females (contrary to males) cluster together with IBERIAN BB females. A mistery then.

@ ROb 3800 BC where? Not the Maykop right? clearly Maykop were not the "humble" people that buried their dead inhouse without ostentation goods as did the Zakubanye culture, but were an offshoot of guys coming from West... that already had the Ostentatious way of life...

You must understand, that most folks in this blog only speak one language and that language is of aDNA . So it will not matter how many references you provide from archaeology,anthropology etc , they will not consider them significant .

You first of all need aDNA from Shulaveri Shomu . Without that neither your thesis can be established or disproved .

@Nirjhar007 Maybe. But until Blog owner cast me out I will keep on making other related points.

Let me give ALL an example. Up until 2 weeks ago, me making the following comment would be receive as baseless. But having found 8000 Balkan farmers with EHG by the buckloads....

One of the odities J. Desideri and Marie bessa found strange is the following: Hungary (Csepel) Bell beakers were closer to Unetice/CWC (eastern CWC) grouping than to BB/CWC in Bohemia. And curiously, Hungary local cultures (Nagyrev, Obebà-Pitvaros and Perjamos) were strangely closer to Bell Beaker/CWC then to Unitece/CWC (eastern).

So, one was to Infer, that those Balkan earlier ( but also contemporaneous and after) cultures (what EHG did they have?) could actually be contributors to Bell beaker/CWC culture in Czech republic hence be a source of the EHG (or part... how much?) you find in Germany BB and also in CWC samples.

I dont know the truth... but who would not brush that immediately if it weren't for the finding of EHG in the balkans 5000 years earlier?

Nirjhar007Yes. We need to wait on the paper. But finding a putative source of the EHG everyone here is talking about as a Yamnaya marker is an raised alarm to yamnayalists (4000 years later).

I am not drawing conclusions. Just that for each new and fresh samples we get tables seem to turn a bit. As it should and as it ALWAYS as happen with new branchs of Science. ALL.That is something you can take to the bank. Always in its infancy more wrong than Right!

Hmm its not clear what they say . Its possible that they had EHG , which will be quite astonishing for some . If there were EHG HG types in Balkans in the Neolithic period , that will turn tables to some degree. Lets see.

@Nirjhar007 Yeah, I am screwd with the shulaveri.Neither the French nor the Japanese seem to get Adna. They get it from goats and cattle... but nope, not from the inhumations next to it!

Actually I need Shulaveri aDNA because I think it would be actually leaning close to Svobodnoe/YAMNAYA even Sintasha and Abashevo... Well They lived in CHG land and some moved to north caucasus. So There you have the CHG in Yamnaya. Then need DNA from Nile delta (5000-4000bc)... see Lots ofvariables. Not easy.

@Nirjhar007,The french must have screw it up.Lyonnet sent lots of samples to be extracted dna in Paris, but it was in 2013 so if nothing came out its because they couldn't.

The Japanese just published a paper regarding Goat DNA in Arukhlo itself, and how those goats actually had come from Eastern Anatolia and not Iran... But don't think they got any Human dna.

Merimda was really important (because morphlogical they were very caucasoid and Armenoid) being so diferent from natufians. That thing is crawling with german teams. But even if they find something they wont publish. Would you?! Saying that the "makers" (not really) of Ancient Egypt were "europeans" (actually not).

I am not drawing conclusions. Just that for each new and fresh samples we get tables seem to turn a bit. As it should and as it ALWAYS as happen with new branchs of Science. ALL.That is something you can take to the bank. Always in its infancy more wrong than Right!

Apart from the fact that you misunderstood the Balkan aDNA abstract, nothing happened.

The Japanese just published a paper regarding Goat DNA in Arukhlo itself, and how those goats actually had come from Eastern Anatolia and not Iran... But don't think they got any Human dna.

Not sure how you missed this, but there are several samples from Chalcolithic Iran, and they look like they came to Iran from the west.

They don't look like the source of southern admixture in Yamnaya though. Wrong Y-DNA and mtDNA.

"Formozov (1965) notes a high density of fortiﬁed Chalcolithic settlements in the northern Caucasus, situated along the Kuban River, which ﬂows into the Sea of Azov at the Taman peninsula. Sites such as SVOBODNOE, MESHOKO, AND ZAMOK BELONG TO THE ZAKUBANYE CULTURE and date to around the SECOND HALF OF THE 5TH MILLENNIUM CAL B.C (Rassamakin 1999;Nekhaev 1992). These were farming villages where large granite cereal grinding stones and ﬂint sickles for cereal cultivation were recovered (Formozov 1965;Korenevskii 2004). Evidence of cereal cultivation near the Ardych-Burunsite was recently published from the Chalcolithic fortiﬁed Svobodnoe site. Macro remains of grain and chaff of Triticum monococcum Triticum dicoccum, and Hordeum vulgare were recovered from the ﬂotation samples (Lebedeva 2011). Furthermore,39% of all tools discovered were attributed to cereal cultivation and processing activities (Nekhaev 1992).

Agriculture in these northern Caucasian sites probably arrived from the southern Caucasus region where large agricultural villages are known to have existed beginning in the early 6th millennium CAL B.C. (Hovsepyan and Willcox 2008; Hovsepyan 2004). It is not clear when the early cultivars started spreading north from present day Georgia and Armenia, reaching the northern portions of the Caucasus. However, the discovery of stratiﬁed Late Mesolithic–Early Neolithic–Bronze Age sites, such as Tsmi (7th to the 3rd millennia CAL B.C )in the northern Caucasus at 1700 masl shows ‘‘the importance of the traverse across the nearby passes in the longue dure´of the Caucasian communication net-work’’ (Rostunov et al.2009: 73) and the possibility that the Neolithic populations of northern Caucasus had contact with the agricultural societies in the south."

and this is called today Shulaveri Shomu.."Agriculture in these northern Caucasian sites probably arrived from the southern Caucasus region where large agricultural villages are known to have existed beginning in the early 6th millennium CAL B.C. (Hovsepyan and Willcox 2008; Hovsepyan 2004). "

@rob,... and did you know that mentioned above Triticum dicoccum is actually a component of SPELT... The same SPELT you see first in Shulaveri, then in Merimda, in early chalcolithic Iberia... and was a trademark of Bell beakers?

I don't think you'll get any arguements that agriculture came to the Kuban via the southern Caucasus But the question is what brought about the rise of the Majkop phenomenon some 1000 years later: refugees from Varna ? Local development ? Or yet again southerners ?

@Rob,My opinion - That Southern Caucasus package (why not call it correctly, SHULAVERI-SHOMU), that had VAST Agriculture and pastoral villages, so loads of people in a very protected and isolated environment by the lesser caucasus mountains and the Caucasus mountain itself, got completely kicked out (or pretty much so) by someone. Hence the question regarding Maykop always was if Maykop had anything to do with them (Shulaveri) with those people that kicked out the Shulaveri and completely dominated the all Caucasus, the middle east and even large parts of Iran, or something coming from the Northwest via balck sea shores. So, all points to the latter. So something Cucuteni-Trypillian, Vinca, Varna also had a story and that story moved into northern and east balck sea. The fact that Maykop never manages even to get really into Lesser Caucasus tells me they were different people from whoever were Kura-araxes main players.

@Davidsky,"Not sure how you missed this, but there are several samples from Chalcolithic Iran, and they look like they came to Iran from the west."

Whatever went from the west to Iran_Chal does not relay at all with Shulaveri. And it looks like Anatolia by mid holocene (say 6000BC) had lots of different people. If they were moving into Europe as Farmers, some others were moving into Anatolia probably as something similar.And a very specific bunch of people, full package highly evolved agriculture, highly evolve pastoral and highly evolved hunters, moved to lesser Caucasus and Kura river and bay. Even once you move south on the Azerbaijan plains into Iran it wasn't "them". And them, did kept lots of horses and apparently didn't eat them (just and oddity).

So, what happen to what could by 5000 bc be tens, upon tens,of thousand of people? Because they Vanished from the region. It was a leveling event of Ashes and their site were not occupied by anyone else. all that at a point when lo and behold agriculture mixed with pastoral starts to appear in several parts of northern Caucasus. - So, Eastern shores of Black sea, Kuban river, Up samara river...

...And who could be the Shulaveri? Well lets sea what those Balkans farmers really look like. Because its near from Bulgaria to Ansueli (where aparently the Shulaveri have the oldest sites) in the shores of the black sea than from Bulgaria to the Azov sea.

@ROb."All this makes sense, so I'm not sure why you're taking the via Alexandria detour for the rest of your thesis"

Because of Humans. And humans have two things: Follow kinship and maintain structural traits as long as are allowed. Non more structural then architectural. The way you Make your house and the way you bury your dead.

Shulaveri were different from all around them. 5 meter huts, 1.5 meter silos and bent walls that unified the houses to make an inner courtyard where they worked and then at the end of day every family went to their own house. - So, the Silo culture (the cultura dos silos)

when they vanished you follow the trail. I found in Tell tsaf (amazingly for a 100-200 year period) at a site of different culture that after them got back to normal rectangular houses. That has a particularity. The oldest copper awl found in the middle east... copper traced to Aruklho itselF!

then I found the same way of life (cultura de silos) in Merimba/el-omari. Then I found it in Iberia as cultura de silos all over South Iberia chalcolithic...

What people don't realize is that Compared to an Europe Route from the steppes to Western Europe, Northern Africa was a highway. Europe was highly densed forrests, rivers upon rivers, upon rivers, mountains upon mountains... and compare to that, north Africa prior the birth of the Sahara was a endless highway of grassland for cattle and humans. Why would they choose to hardest possible way if they had an open space!?